DOGROSES (ROSA SECT. CANINAE L.), OLD MEDICINAL PLANTS … · 2020-07-18 · MEDICINAL AND AROMATIC...

20
MEDICINAL AND AROMATIC PLANTS OF THE WORLD – Vol. I - Dogroses (Rosa Sect. Caninae L.), Old Medicinal Plants With A Large Potential – A Review - Hilde Nybom and Gun Werlemark ©Encyclopedia of Life Support Systems (EOLSS) DOGROSES (ROSA SECT. CANINAE L.), OLD MEDICINAL PLANTS WITH A LARGE POTENTIAL – A REVIEW Hilde Nybom and Gun Werlemark Department for Plant Beeding and Biotechnology-Balsgård, Swedish University of Agricultural Sciences, Fjälkestadsvägen 459, SE 29194 Kristianstad, Sweden Keywords: pentaploids, odd meiosis, complex taxonomy, high antioxidant activity, polyphenols, carotenoids, vitamin C, B and E, fatty acids, anti-inflammatory and antinociceptive substances, anti-diabetic, cosmetics, seed-propagated Contents 1. Introduction 2. Taxonomy 3. Cytology and reproduction 4. Genetics and plant breeding 5. Health-related properties 6. Chemical contents 6.1. Rosehips 6.2. Seeds 7. Processing 8. Utilization as food and medicine 9. Commercial rosehip production 9.1. Propagation 9.2. Planting 9.3. Pruning 9.4. Harvesting 9.5. Post Harvest Handling 10. Global production and trade Glossary Bibliography Biographical Sketches Summary Dogroses, i.e. species belonging to Rosa sect. Caninae L., grow wild along roadsides and in disturbed areas in temperate regions, and have also been cultivated in some countries to a lesser extent. However, the newly acquired interest in food products with health-promoting substances may lead to new plantations. Rosehips from dogroses have been shown to contain very high levels of antioxidant compounds, mainly polyphenols, but also carotenoids and the vitamins B, C and E. Several different studies have shown rosehip extract to have anti-inflammatory, antinociceptive, anti-diabetic and anti- mutagenic effects. It has also been shown to inhibit certain cancer cell proliferation in vitro. The most well-performed studies so far concern the positive effect on patients with osteoarthritis, as well as effects on various stomach problems. Rose hip extract has also been shown to have an inhibitory effect on body weight gain, and to reduce the glucose level in the blood stream making it useful as a treatment against obesity. The

Transcript of DOGROSES (ROSA SECT. CANINAE L.), OLD MEDICINAL PLANTS … · 2020-07-18 · MEDICINAL AND AROMATIC...

Page 1: DOGROSES (ROSA SECT. CANINAE L.), OLD MEDICINAL PLANTS … · 2020-07-18 · MEDICINAL AND AROMATIC PLANTS OF THE WORLD – Vol. I - Dogroses (Rosa Sect. Caninae L.), Old Medicinal

MEDICINAL AND AROMATIC PLANTS OF THE WORLD – Vol. I - Dogroses (Rosa Sect. Caninae L.), Old Medicinal Plants With A Large Potential – A Review - Hilde Nybom and Gun Werlemark

©Encyclopedia of Life Support Systems (EOLSS)

DOGROSES (ROSA SECT. CANINAE L.), OLD MEDICINAL

PLANTS WITH A LARGE POTENTIAL – A REVIEW

Hilde Nybom and Gun Werlemark

Department for Plant Beeding and Biotechnology-Balsgård, Swedish University of

Agricultural Sciences, Fjälkestadsvägen 459, SE 29194 Kristianstad, Sweden

Keywords: pentaploids, odd meiosis, complex taxonomy, high antioxidant activity,

polyphenols, carotenoids, vitamin C, B and E, fatty acids, anti-inflammatory and

antinociceptive substances, anti-diabetic, cosmetics, seed-propagated

Contents

1. Introduction

2. Taxonomy

3. Cytology and reproduction

4. Genetics and plant breeding

5. Health-related properties

6. Chemical contents

6.1. Rosehips

6.2. Seeds

7. Processing

8. Utilization as food and medicine

9. Commercial rosehip production

9.1. Propagation

9.2. Planting

9.3. Pruning

9.4. Harvesting

9.5. Post Harvest Handling

10. Global production and trade

Glossary

Bibliography

Biographical Sketches

Summary

Dogroses, i.e. species belonging to Rosa sect. Caninae L., grow wild along roadsides

and in disturbed areas in temperate regions, and have also been cultivated in some

countries to a lesser extent. However, the newly acquired interest in food products with

health-promoting substances may lead to new plantations. Rosehips from dogroses have

been shown to contain very high levels of antioxidant compounds, mainly polyphenols,

but also carotenoids and the vitamins B, C and E. Several different studies have shown

rosehip extract to have anti-inflammatory, antinociceptive, anti-diabetic and anti-

mutagenic effects. It has also been shown to inhibit certain cancer cell proliferation in

vitro. The most well-performed studies so far concern the positive effect on patients

with osteoarthritis, as well as effects on various stomach problems. Rose hip extract has

also been shown to have an inhibitory effect on body weight gain, and to reduce the

glucose level in the blood stream making it useful as a treatment against obesity. The

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©Encyclopedia of Life Support Systems (EOLSS)

seeds contain high levels of unsaturated fatty acids, which can be used for skin

treatment and cosmetics. The amount of these bioactive compounds varies with

genotype and environment. The dogroses can be seed-propagated, but as the

germination is slow, cuttings are preferred. The rosehips are mainly harvested by hand,

and should be picked when almost ripe, and then dried or frozen as soon as possible, to

ensure high contents of bioactive compounds. Presently, production of dogrose rosehips

is increasing due to the big interest from health-conscious consumers.

1. Introduction

The well-known genus Rosa in the family Rosaceae contains numerous species of

commercial interest. Several wild rose species like Rosa moschata, R. wichurana, R.

multiflora, R. damascena, R. gallica, R. chinensis, R. gigantea, and R. foetida have

contributed to the highly appreciated and economically important ornamental rose

cultivars, and in the case of R. damascena also to the rose oil industry.

Other rose species have been less noted by the rose-growing industry but some have the

potential to become very important due to their fruits, the rosehips. Commercial rosehip

production involves several different rose species. One important group is the dogroses,

all of which belong to the section Caninae. These are long-lived woody perennials,

growing in woodland margins and disturbed habitats such as roadsides and open

pastures. Dogroses are sometimes planted as ornamentals, especially in public areas

where they are appreciated for their rustic appearance and hardiness. Several dogrose

species have also been used as rootstocks for the budding of ornamental roses, but other

rootstocks with superior characteristics are now being used to an increasing extent. In

the middle ages, dogroses were however cultivated, especially at monasteries, for use as

valuable medicinal plants. Almost all parts of the plants were used: rosehips, seeds,

petals, leaves and roots. During the last two decades, a renewed interest in the dogrose

species have emerged since modern science have shown that rosehips contain valuable

compounds that can be used as functional food or nutraceuticals.

2. Taxonomy

Rose taxonomy is quite complicated and there have been several attempts to improve

the classification of roses through the years. For lack of a commonly accepted

alternative, most people still use the system of Rehder (1940), updated by Wissemann

(2003), which comprises four subgenera: Hulthemia (1 species), Hesperhodos (two

species), Plathyrhodon (one species) and Rosa (approx. 180 species). Subgenus Rosa is

subdivided into 10 different sections, and the largest of these sections is sect. Caninae,

dogroses, with approx. 50 species. The other sections are Pimpinellifoliae (15 species),

Rosa (only one proper species but several hybrid taxa have been given species rank),

Carolinae (possibly five species), Cinnamomeae (about 80 species), Synstylae (about 25

species), Indicae (three species), Banksianae (possibly two species), Laevigatae (one

species) and Bracteatae (one or two species).

Within the genus Rosa, section Caninae has proved to be the most difficult group. At

the beginning of the last century, the name-giving was at its peak when the Swedish

rhodologist Almqvist (1919) described more than 350 dogrose species on one island

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alone in the Stockholm archipelago! Today, there is a more restricted view on what

entity is entitled species rank, and the number of recognized dogrose species presently

hoovers around 50 (Wissemann 2003). Up to six subsections are generally recognized;

subsect. Trachyphyllae with R. jundzillii as the only species, subsect. Rubrifoliae with

R. glauca (syn. R. rubrifolia) as the only species, subsect. Vestitae with, e.g., R. mollis,

R. pseudoscabriuscula, R. sherardii, R. tomentosa and R. villosa (syn. R. pomifera),

subsect. Rubigineae with, e.g., R. agrestis, R. inodora, R. micrantha., and R. rubiginosa,

subsect. Tomentellae with R. tomentella and R. abietina, and subsect. Caninae with,

e.g., R. canina, R. corymbifera, R. dumalis, R. montana, R. stylosa, R. subcanina and R.

subcollina (Wissemann 2003).

Traditionally, dogrose species have been defined according to morphological characters

like leaflet shape, hip shape, length of pedicel, presence or absence of glandular hairs,

shape of prickles, shape of style head orifice, leaf pubescence, and plant shape.

Although some discontinuities are noted, these characters have often been insufficient

for unambiguous classification of plant material, even when this material has been

grown in comparative garden trials (Nybom et al. 1996, Olsson et al. 2000, De Cock et

al. 2008).

Figure 1. Rosa rubiginosa in full bloom

In recent years, morphology-based information has often been complemented with

DNA-based data for plant classification studies. In dogroses, many taxa have, however,

turned out to be even more difficult to distinguish when analysed with DNA markers

compared to when analysed with morphological characters. Samples from closely

related species (belonging to the same subsection) have often turned out to overlap

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©Encyclopedia of Life Support Systems (EOLSS)

completely when analysed with DNA markers like RAPD and AFLP (Olsson et al.

2000, Atienza et al. 2005, de Cock et al. 2008, Koopman et al. 2008). By contrast, the

same studies have usually reported some level of discontinuity between at least the three

major subsections: Vestitae, Rubigineae and Caninae suggesting that these constitute

biologically valid and identifiable taxa.

In this report, we present a substantial amount of the research so far published on rose

species suitable for rosehip production, with a major emphasis on the dogrose species.

3. Cytology and Reproduction

In the genus Rosa, the basic chromosome number x = 7. Diploid rose species thus have

two sets of chromosomes, 2n = 14. In meiosis, these 14 chromosomes pair up to form

seven bivalents as expected, resulting in regular biparental inheritance. Many rose

species as well as almost all ornamental cultivars are instead tetraploid, with 2n = 28.

Also these have a regular bivalent formation.

Figure 2. Canina meiosis. In the female meiosis, 14 chromosomes form 7 bivalents

whereas the remaining chromosomes occur as univalents and gather towards one side of

the cell. The bivalent chromosomes pair and recombine, and after the separation and

second division, two viable cells are formed with 7 chromosomes from the bivalent

formation and all the univalents, and two inviable cells with only the 7 chromosomes

from the bivalent formation. At the male side, 7 bivalents are again formed, but this

time the univalents start to separate into chromatids already in the first metaphase. After

the second division, most of the univalents are lost, or occur as chromatids or

chromosome parts in some of the pollen cells. Viable pollen cells contain only the 7

chromosomes from the bivalent formation; pollen viability in the section Caninae is

only 20–35%.

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There is, however, one notable exception in the genus. Thus, all dogroses (sect.

Caninae) are characterized by the peculiar canina meiosis described over 80 years ago

(Täckholm, 1922) and verified in several, more recent publications (e.g. Lim et al.,

2005). The dogroses are usually pentaploid although some tetraploids and hexaploids

also occur. Regardless of ploidy level, only seven bivalents are formed in the first

meiotic division. The remaining chromosomes occur exclusively as univalents. These

univalents are not included in viable pollen grains, which contain only the seven divided

bivalent chromosomes. In contrast, all the univalents are transmitted to one of the

daughter cells in the female meiosis, and are finally included in the viable egg cells,

which therefore contain 21, 28 or 35 chromosomes, depending on the ploidy level. The

resulting seedlings obtain the full chromosome number but only 15–25% of these

chromosomes are inherited from the pollen parent, whereas the remainder are obtained

from the seed parent. This means that the offspring usually is much more similar to the

maternal parent (matroclinal inheritance) than to the paternal parent. In addition, results

from morphological evaluation of spontaneous or experimentally derived dogrose

progenies has led some scientists, e.g., Gustafsson (1937, 1944) and Kroon and Zeilinga

(1974) to suggest an apomictic seed production, i.e. seeds being produced without prior

fertiliztion of the egg cell. Recently, up to 5 or 10% apomictic seed set has also been

demonstrated with DNA markers in som dogrose species (Werlemark et al., 1999;

Werlemark, 2000; Werlemark and Nybom, 2001; Nybom et al., 2004a, 2006).

Diploid rose species are usually selfsterile which means that they have to be

crosspollinated in order to set seed. By contrast, polyploid rose species, including the

dogroses, are usually self-fertile and can therefore set seed after selfing. Breeding

behaviour affects the amount and partitioning of genetic variation in wild populations,

and in the offspring of plants used in breeding programmes.

4. Genetics and Plant Breeding

Presently, precise information on character inheritance has been developed mostly for

ornamental rose cultivars in traits that have little or no relevance for rosehip production,

like flower morphology, colour and fragrance, moss character, dwarf phenotype, stem

prickles, petiole prickles, foliage glossiness and recurrent blooming (Debener, 2003;

Byrne 2009). Although highly desirable in ornamentals, the recessively inherited

recurrent blooming trait is a serious draw-back in roses grown for hip production,

especially in machine-harvested orchards since repeated harvesting is required. Lack of

prickles on the stems would improve manual orchard work but could also render the

plants more vulnerable to grazing animals in orchards with inadequate fencing. When

prickles are present, their number and size are quantitatively inherited (Lal et al., 1082).

The climbing growth type has been reported to be dominant over the non-climbing

growth type in ornamental roses (Morey, 1954). Interestingly, the arching growth type

in dogroses has also been reported to be dominant over the more desirable erect (and

non-climbing) growth type (Wissemann et al., 2006). Other growth type-related traits

studied in ornamental roses like length of the flowering stems and plant height, as well

as branch number and plant spread, are under polygenic control with a high heritability

(Lal et al., 1982). Similarly, most of the vigour-related traits so far investigated in

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connection with ornamental rose breeding, appear to be under polygenic control (Byrne

2009).

Analyses of resistance against fungal diseases in ornamental roses and some diploid

wild species, have demonstrated both polygenically controlled, horizontal resistance and

vertical, race-specific resistance conferred by major genes, e.g., Rdr1 and Rbs providing

resistance against blackspot, Marssonina rosae (Kaufmann et al., 2003; Yan et al.,

2005b), and Rpp1 and Rpm providing resistance against powdery mildew (Linde et al.,

2004; Zhang, 2003). In dogroses, both blackspot and powdery mildew instead appear to

be under polygenic control since at least some symptoms can be found on all

investigated species so far (Carlson-Nilsson and Uggla, 2005; Uggla and Carlson-

Nilsson, 2005; Schwer et al., 2007). Similarly, rust and Sphaceloma-leafspot occur in

all dogrose species evaluated although at variable levels (Schwer et al., 2007). Boerema

(1963) reports that some dogrose species like R. canina, R. tomentosa, R. villosa and R.

rubiginosa are susceptible to Septoria-leafspot, whereas other species lack symptoms

altogether. In a Swedish study, R. rubiginosa had no symptoms although other species

in the same field were heavily infested, suggesting a genetically controlled, perhaps

monogenic resistance (Schwer et al., 2007).

Figure 3. The fungal disease blackspot (Diplocarpon rosae perfect stage, or Marssonina

rosae imperfect stage)

Figure 4. The fungal disease leafspot (Sphaceloma rosarum) also known as anthracnose

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So far, little if any directed plant breeding has been undertaken for increasing contents

of bioactive compounds in rosehips. Considerable levels of inter- and intraspecific

variation has, however, been reported for various compounds like ascorbic acid,

lycopene, minerals, phenolic compounds and total antioxidant activity (Celik et al.

2009; Günes 2010; Turkben et al. 2010a). These results suggest that there is a

considerable potential for developing new rose hip cultivars with more desirable

chemical contents.

5. Health-Related Properties

Nowadays, people are becoming increasingly concerned about issues related to food

and health. Fruit and berries are appreciated for their often high contents of health-

promoting compounds. In this context, dogroses have been regarded as important

medicinal plants since the middle ages. Rosehips, as well as petals, leaves and seeds,

have been used for all sorts of concoctions and tonics to treat a wide range of ailments.

Therapeutic activities have thus been described against the common cold, gastro-

intestinal disorders and gastric ulcers, and different kinds of inflammatory disorders,

e.g., arthritis. Uses as a laxative and a diuretic have also been reported.

The mechanisms responsible for possible medicinal effects of rosehips have, for the

most part, not yet been discovered. Chemical analyses of bioactive compounds have

been undertaken as a step in revealing health-promoting activities. Rosehips from

dogroses have thus been shown to contain very high levels of ascorbic acid, and to have

the highest percentage of total antioxidants among a large set of different fruit and berry

crops (Halvorsen et al. 2002). Lycopene levels equal those found in tomatoes (Böhm et

al. 2003), and there is also a high level of folate (vitamin B) (Strålsjö et al. 2003).

Rosehip seeds are very rich in unsaturated fatty acids (Szentmihályi et al. 2002). Many

medicinal properties have thus been reported for rosehip extracts, e.g. growth inhibition

of certain cancer cells (Olsson et al. 2004), decreased activity of reactive oxygen species

(ROS) in rat colons (Håkansson et al. 2006), and an anti-inflammatory response when

consumed by patients with osteoarthritis (Winther et al. 2005).

6. Chemical Contents

Unfortunately, it is almost impossible to compare different reports on the bioactive

compounds in dogroses. To begin with, very few of the authors of these studies are

aware of the taxonomic complexity within section Caninae. Instead, the studied

rosehips are often said to have been collected from R. canina even though they may

actually have been collected from a different species within the section, or even from

another section. Sometimes the species is not mentioned at all; perhaps the extraction

was made on material bought in another country or picked in the backyard. Also,

several studies show that there is variation in concentration of bioactive compounds

between genotypes within one species, even if they grow in the same place (Uggla et al.

2003). The composition of bioactive compounds is also affected by the ripening stage of

the rosehips at harvest (Guimaraes et al. 2010). Similarly, blooming stage of harvested

R. damascena flowers affect the amount and quality of the industrial rose oil obtained

(Baydar & Baydar 2004).

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The chemical contents of rosehips are also dependent on various environmental factors

(Demir & Özcan 2000, Celik et al. 2009, Ghazghazi et al. 2010). Differences in

composition between years depending upon precipitation, temperatures, insolation etc

have thus been reported (Kovacs & Toth 2000, Strålsjö et al. 2003, Uggla et al. 2003,

Kähkönen et al. 2001, Ross & Kasum 2002). The latitude of the location also appears to

affect the biochemical composition (Keles 2007).

Similarly, the amount of polyphenols in the leaves is affected by, e.g., gall insect

infestations (Coruh & Ercisli 2010). Chemical contents are also dependent upon the

post-harvest procedures, e.g, on how long time it takes before the rosehips are frozen or

dried. Traditionally, rosehips are put out in the sun to dry, which can take several days

during which the contents may deteriorate. Finally, extraction methods are also

important for measuring of chemical contents

All papers cited in our study have been published in peer-reviewed journals. Still,

reviewers have not always noticed if the authors mentioned whether they studied dry or

fresh material; concentrations are of course higher per gram in dry material compared to

in fresh material. The authors also report concentrations in different ways, e.g. g/kg FW,

µg/g DW, mg GAE/g DW, mmol/g FW etc, making it difficult to compare the different

results.

Also, some authors do not mention whether they have used whole rosehips with seeds

or rosehips without seed. Dogrose-derived rosehips contain 25–40 seeds/hip, which is

approximately ¼ of the total weight of the hip. These seeds contain over 77%

polyunsaturated fatty acids (PUFA), which are very susceptible to chemical reactions

and can cause rapid deterioration of other compounds (Concha et al. 2006).

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Glossary

AFLP : Amplified fragment length polymorphism

α-tocopherol : Vitamin E

All-trans-retinoic

acid :

ATRA or tretinoin. A natural pre-cursor of vitamin A used to

treat different skin problems and in cosmetics, found in rosehip

seeds

Anaerobic

conditions :

Without oxygen

Antinociceptive : Reducing sensitivity to painful stimuli

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Antioxidant : A molecule capable of inhibiting oxidation of other molecules

and prevent ROS damage by scavenging the free radicals.

Apomixis : The ability to set seed without prior fertiization, offspring

becomes identical to mother plant

Ascorbic acid : Vitamin C

Carotenoids : Antioxidants that can be divided into two groups, xanthophylls

and carotenes. Within the carotenes there are α-carotene, β-

carotene (precursor to vitamin A) and lycopene.

Chemotaxis : Movement of cells or organisms along a chemical concentration

gradient either towards or away from the simulus.

COX : Cyclooxygenase enzymes responsible for inflammation and pain.

DW : Dry weight

Ellagic acid : A polyphenolic tannin compound, claimed to have beneficial

effects against cancer and other medicinal problems. High levels

have been found in raspberries, strawberries and rosehips

Flavonoids : Type of polyphenols, consisting of six major subclasses:

flavones, flavonols, flavanones, cathechins, anthocyanidins and

isoflavones. The flavonoids act as antioxidants and inhibits the

oxidation of low density lipoproteins and prevent aggregation of

platelets.

Folate : Vitamin B9

Functional food : A food where a new ingredient/s has been added or where an

existing ingredient has been increased, and the new product has a

new function, often related to health promotion or disease

prevention.

FW : Fresh weight

GAE : Gallic acid equivalent

Gastrointestinal

tract :

The stomach and intestine

Horizontal

resistance :

Many genes are involved in the resistance, usually provides

unspecific, durable resistance

Interspecific

variation :

Variation between the species

Intraspecific

variation :

Variation within the species

In vitro : The biological experiment is conducted outside the organism

In vivo : The biological experiment is conducted inside the living

organism

Linoleic acid : Omega-6, an essential fatty acid in rosehip seeds

Linolenic acid : Omega-3, an essential fatty acid in rosehip seeds

Lipogenic : Producing, forming or caused by fat

Matroclinal

inheritance :

The genetic material in the offspring is inherited mainly from the

seed parent

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Meiosis : Chromosome division, takes place immediately before the

formation of egg cells and pollen cells, usually results in a 50%

reduction of the chromosome number.

Nutraceuticals : “a product isolated or purified from foods that is generally sold in

medicinal forms not usually associated with food. A nutraceutical

is demonstrated to have a physiological benefit or provide

protection against chronic disease” Health Canada definition. The

products may range from isolated nutrients, dietary supplements

and specific diets to herbal products and processed foods.

Osteoarthritis : A progressive disorder of the joints caused by gradual loss of

cartilage. One of the most common causes of disability especially

in people over 50.

Ploidy level : Refers to the number of chromosomes. In the genus Rosa, the

basic chromosome number is 7, diploid species have 14

chromosomes, triploid have 21, tetraploid have 28, pentaploid

have 35 and hexaploid have 42.

Pro-oxidant : Can induce oxidative stress either through creating ROS or

inhabiting antioxidant systems

RAPD : Random amplified polymorphic DNA

RNS : Reactive nitrogen species

ROS : Reactive oxidative species, forms naturally in the body as a

byproduct of normal metabolism so called free radicals.

However, in stress situations e.g. sickness, smoking, radiation,

ROS levels can increase to high levels and cause significant

damage to cell structures.

Selfing : Self-pollination, generally produces very homogenous offspring

Supercritical CO2 : Fluid state of CO2 used as solvent with low toxicity and

environmental impact

Unsaturated fatty

acids :

A fatty acid with at least one double bond within the fatty acid

chain and thus an elimination of one hydrogen atom, whereas

saturated fatty acids do not have any double or triple bonds. An

unsaturated fatty acid contains less energy (calories) than a

saturated fatty acid. Diets high in polyunsaturated fatty acids and

low in saturated fatty acids have been correlated with low serum

cholesterol levels in some studies.

Vertical resistance : A single gene is responsible for the resistance, often provides

race-specific resistance, prone to „resistanc breaking‟

Bibliography

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©Encyclopedia of Life Support Systems (EOLSS)

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©Encyclopedia of Life Support Systems (EOLSS)

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[Antioxidant activity was studied in rosehips from Polish roses].

Nowak R., Gawlik-Dziki U. (2007) Polyphenols of Rosa L. leaves extracts and their radical scavenging

activity. Zeitschrift Naturforschung, 62, 32-38 [Polyphenolic contents were determined in leaves from 17

Polish Rosa species].

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Caninae, Rosaceae). Acta Universitatis Upsaliensis Symbolae botanicae Upsalienses 31, 59-68

[Systematics in Swedish dogroses based on morphometrics].

Nybom H., Esselink G.D., Werlemark G., Vosman B. (2004) Microsatellite DNA marker inheritance

indicates preferential pairing between highly homologous genomes in polyploidy and hemisexual

dogroses Rosa L. sect. Caninae. Heredity 92, 139-150 [Analysis of quantitative peak differences allowed

a tentative estimation of chromosome transmittal and allelic configuration in individual dogrose plants].

Nybom H., Esselink G.D., Werlemark G., Leus K., Vosman B. (2006) Unique genomic configuration

revealed by microsatellite DNA in polyploidy dogroses, Rosa sect. Caninae. Journal of Evolutionary

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Biology 19, 635-648 [The allopolyploid complex of dogroses was studied; the maternally transmitted

univalent chromosomes reflect taxonomic distances to a higher degree than the biparentally inherited

bivalent chromosomes].

Ochoa M.R., Kesseler A.G., Pirone N., Marquez A., de Michelis A. (2002) Shrinkage during convective

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Caninae, Rosaceae) assessed by RAPDs and elliptic Fourier analysis of leaflet shape. Systematic Botany

25, 511-521 [Taxonomic studies of Nordic dogroses using DNA markers and morphometrics].

Olsson M.E., Gustavsson K.-E., Andersson S., Nilsson Å., Duan R.-D. (2004) Inhibition of cancer cell

proliferation in vitro by fruit and berry extracts and correlations with antioxidant levels. Journal of

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cell proliferation of colon cancer cells were investigated].

Olsson M.E., Andersson S., Werlemark G., Uggla M., Gustavsson K.E. (2005) Carotenoids and phenolics

in rose hips. Acta Horticulturae 690, 249-252 [Fruits from ten different Rosa species were analysed for

their contents of carotenoids and phenolics].

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of Rosa canina fruits. Pharmacognosy Magazine 5, 309-315 [Ethanol extracts of R. canina fruits were

screened for their antioxidant, hypoglycaemic and antidiabetic activities. The findings support the

traditional use of R. canina fruits as a folk remedy of diabetes in Turkey].

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chloro-ascorbid acid: potential anticancer drugs. Neoplasma 44, 101-107 [The influence of ascorbic acid

on the growth of various human cell lines was examined. Inhibitory effect was confirmed].

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tocopherol transfer protein. Nature Genetics 9, 141-145 [Ataxia with isolated vitamin E deficiency

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protectively against neurological damage in this disease].

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vitro. Three cases are presented].

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of ascorbic acid concentration during dehydration of rosehip (Rosa eglanteria) fruits. American Journal

of Food Technology 2, 377-387 [A study to ascertain the degradation of ascorbic acid as a function of

drying temperature].

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bioproducts. Asia-Pacific Journal of Chemical Engineering 2, 20-29 [Different techniques for drying of

biological material].

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system for the genus Rosa; much criticized but still the basis for almost all systematic Rosa papers in the

last 50-60 years].

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bushes].

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Ross J.A., Kasum C.M. (2002) Dietary flavonoids: bioavailability, metabolic effects, and safety. Annual

Review of Nutrition 22, 19-34 [Review of flavonoids, biochemistry and possible health effects].

Salminen J.-P., Karonen M., Lempa K., Liimatainen J., Sinkkonen J., Lukkarinen M., Pihlaja K. (2005)

Characterisation of proanthocyanidin aglycones and glycosides from rose hips by high-performance

liquid chromatography-mass spectrometry, and their rapid quantification together with vitamin C. Journal

of Chromatography A 1077, 170-180 [15 proanthocyanidin aglycones and 19 glycosides were found in

dogrose rosehips, as well as high levels of vitamin C].

Sanderson K., Fillmore S. (2010) Evaluation of native rose selections for rose hip production in Prince

Edward Island. International Journal of Fruit Science 10, 379-389 [A study to determine the potential for

commercial production of rosehips on Prince Edward Island, Canada].

Schwer C.S., Carlson-Nilsson U., Uggla M., Werlemark G., Nybom H. (2007) Impact of foliar fungi on

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some species of dogroses in Sweden].

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ssp.) after drying. Journal of Agricultural and Food Chemistry 51, 4291-4295 [Folate concentrations in

rosehips, and commercial products and factors affecting folate retention during drying were investigated].

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methods of valuable rosehip oil].

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2309-2318 [Chemical content in rosehips from different Rosa species were analysed, before and after

processing].

Turkben C., Uylaser V., Incendayi B., Celikkol I. (2010b) Effects of different maturity periods and

processes on nutritional components of rose hip (Rosa canina L.). Journal of Food Agriculture &

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naturally and other processed traditionally. Contents were analysed].

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during ripening. PhD thesis, Acta Universitatis Agriculturae Sueciae, Agraria 480 [Determination of

optimal harvest time of R. spinosissima using fruit detachment force and quality parameters].

Uggla M., Gao X., Werlemark G. (2003) Variation among and within dogrose taxa (Rosa sect. Caninae)

in fruit weight, percentages of fruit flesh and dry matter, and vitamin C content. Acta Agriculturae

Scandinavica, Sect B, Soil Plant Science 53, 147-155 [Variation among dogrose taxa studied with

different quality parameters].

Uggla M., Carlson-Nilsson B.U. (2005) Screening of fungal diseases in offspring from crosses between

Rosa sections Caninae and Cinnamomae. Scientia Horticulturae 104, 493-504 [Variation among

hybridogenous Rosa plants in tolerance towards fungal infection, indicates the possibility to select for

resistance to serious diseases].

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Horticulturae 690, 83-89 [Description of procedures involved in commercial field-based rosehip

cultivation in Sweden].

Uggla M., Gustavsson K.E., Olsson M.E., Nybom H. (2005) Changes in color and sugar content in

rosehips (Rosa dumalis L. and R. rubiginosa L.) during ripening. Journal of Horticultural Science and

Biotechnology 80, 204-208 [The color of rosehips changes with maturation, and can be used as an

indicator of sugar content and optimal harvesting time].

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Ullah M.F., Khan H.Y., Zubair H., Shamim U., Hadi S.M. (2011) The antioxidant ascorbic acid mobilizes

nuclear copper leading to a prooxidant breakage of cellular DNA: implications for chemotherapeutic

action against cancer. Cancer Chemotherapy and Pharmacology 67, 103-110 [Ascorbic acid has

previously ben shown to be cytotoxic to certain cancer cells. This study proposes the cause to be the

elevated copper levels in the cancer cells].

Umezu T. (1999) Anticonflict effects of plant-derived essential oils. Pharmacology, Biochemistry and

Behaviour 64, 35-40 [A study on mice showed that rose oil can be used to obtain anticonflict effects].

Upston J.M., Kritharides L., Stocker R. (2003) The role of vitamin E in atherosclerosis. Progress in Lipid

Research 42, 405-422 [There is a balance between vitamin E and other antioxidants that is crucial for in

vivo antioxidant protection in atherosclerosis, but vitamin E is not deficient in atherosclerosis lesions]

Warholm O., Skaar S., Hedman E., Mølmen H.M., Eik L. (2003) The effects of a standardized herbal

remedy made from a subtype of Rosa canina in patients with osteoarthritis: A double-blind, randomized,

placebo-controlled clinical trial. Current Therapeutic Research 64, 21-31 [Standardized rosehip powder

was shown to reduce symptoms of osteoarthritis in patients with a diagnosis of osteoarthritis in either hip

or knee].

Werlemark G. (2000) Evidence of apomixis in hemisexual dogrose species, Rosa sect Caninae. Sexual

Plant Reproduction 12, 353-359 (Apomixis, i.e. seed set without fertilization, is shown to exist in at least

some dogrose species].

Werlemark G., Uggla M., Nybom H. (1999) Highly skewed distribution of morphological and RAPD

markers in a pair of reciprocal crosses between hemisexual dogrose species, Rosa sect. Caninae.

Theoretical and Applied Genetics 98, 557-563. [The uneven distribution of inherited traits in dogroses is

shown with DNA markers and morphological characters].

Werlemark G., Nybom H. (2001) Skewed distribution of morphological character scores and molecular

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distribution of markers in dogroses is shown in the offspring from three interspecific crosses].

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seedling emergence in dogroses, Rosa sect. Caninae L. Acta Agriculturae Scandinavica, Sect B, Soil

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Winther K., Apel K., Thamsborg G. (2005) A powder made from seeds and shells of a rose-hip

subspecies (Rosa canina) reduces symptoms of knee and hip osteoarthritis: a randomized double-blind,

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currant and rosehip was examined].

Biographical Sketches

Gun Werlemark received her Ph D in plant breeding at the Swedish University of Agricultural Sciences

(SLU) in 2001. She has since worked at the Department of Plant Breeding and Biotechnology, Balsgård,

SLU, as a researcher. Her main interest has been the genetics of dogroses studied with molecular,

morphological and phytochemical markers. In addition, she has worked with the application of different

types of DNA markers in several other horticulturally important plants like Rubus (blackberries and

raspberries) and Nymphaea (water lilies) Recently she has also extended her interest to the genetics and

phytochemistry of the medicinally important bilberries (Vaccinium myrtillus). She is the author/co-author

of some 20 scientific papers.

Hilde Nybom, received her Ph D in plant systematics at Lund University in 1987. She is a professor at

the Department of Plant Breeding and Biotechnology, Balsgård, SLU. She is the author/co-author of

approximately 100 scientific papers and of two textbooks on DNA fingerprinting in plants. She received

the 'Golden Apple' award in 2006 for research and breeding of Nordic fruit and berries, and the

‟Environmental-Medicinal prize‟ in 2010 for research on low-allergenic apple cultivars. Her present

research interests concern plant systematics and population genetics, fruit and berry genetics and

breeding, rose genetics, and development of horticultural crops for industrial and/or medicinal use.